Discussion forum

Integrating bottom-up internalist views ofemotional feelings with top-down externalistviews: Might brain affective changes constitutereward and punishment effects within animalbrains?

Jaak Panksepp*

Baily Endowed Chair of Animal Well-Being Science, Department of Integrative Physiology and Neuroscience, Collegeof Veterinary Medicine, Washington State University, Pullman, WA, USA

In “Emotion and DecisionMaking Explained” Edmund Rolls sharesan exceptionally well-detailed, deeply scholarly work thatprobes the broad landscape of animal and human emotions,as well as body regulatory processes such as hunger and thirstand sensory affects, especially taste. Rolls’s extensive syn-thesis into a cognitive (higher order thought) theory of con-sciousness is an impressive integration of an enormousamount of empirical work from both human and animalneurosciences, culminating in a cognitive view of mental life,where intrinsic affective (survival) values encoded in brains arenot as critical for feelings as higher-order thoughts. It is amust read for anyone interested in the details of how the brainmay construct mental experiences. His cross-species neuro-science synthesis is contextualized in both, a traditionalbehavioristic (reinforcement) as well as cognitive appraisaland social-constructivist frameworks, concluding with a so-phisticated computational theory of mind/consciousness.

In this superlative synthesis, Rolls discusses issues of first-rate importance for understanding brain functions andhigher-order consciousness. I will only focus on the one largepotential flaw in his arguments: Perhaps he unfairly dismissesthe existence of a more primal form of consciousness, namelyraw (unreflected-upon) affective phenomenal experiences inother animals? He may have fallen into this potential trap

because he is developing a top-down view of the consciousmind, asserting that the internally experienced aspects ofbrain functions are based almost exclusively on the self-reflective capacities afforded by language-type (syntactic)brain functions (pp. 487e488), a view that is surely debatable,and one with which I disagree, partly because it seems soinconsistent with graded evolutionary progressions in all life-sustaining functions. Rolls’s sophisticated top-down view ofthe experienced mind, with the central assertion thatconscious experience in humans is based on syntacticthoughts (implicit “language”?), does not dwell on the alter-native evidence-based possibility that the capacity to haveraw affective experiences is largely a bottom-up, evolved-adaptive aspect ofmind, based on intrinsic biological “values”,shared by many creaturesda neurodynamic survival-enhancing process that may not require sophisticated top-down reflective aspects (typically called “awareness”done ofthe most slippery terms in consciousness studies) in order tobe experienced.

Admittedly, what I will focus on is only a small fraction ofhis impressive literature synthesis and theory development,but it is an issue ofmonumental importance. Regrettably suchtopics have the flickering quality of Necker Cubes, dependingon whether one takes a top-down self-reflective perspectives

* Department of Integrative Physiology and Neuroscience, College of Veterinary Medicine, Washington State University, PO Box 647620,Pullman, WA 99164-7620, USA.

E-mail address: jpanksepp@vetmed.wsu.edu.

Available online at www.sciencedirect.com

ScienceDirectJournal homepage: www.elsevier.com/locate/cortex

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Please cite this article in press as: Panksepp, J., Integrating bottom-up internalist views of emotional feelings with top-downexternalist views: Might brain affective changes constitute reward and punishment effects within animal brains?, Cortex(2014), http://dx.doi.org/10.1016/j.cortex.2014.04.015

on themental apparatus, as Rolls does, or bottom-up, evolved-valuative views that I prefer. Clearly the two can be synthe-sized, so I would ask Rolls to reconsider the possibility that araw, non-reflective “affective consciousness”dan intrinsic“feeling” property of reverberating brain networks in at leastmammalian brainsdevolved much earlier than our capacityto syntactically reflect on our many raw experiences. Indeed,from my perspective, perhaps the process of “reinforcement”is how the primary-process (unconditional) circuits for affec-tive values, arising from deep-subcortical networks, allowsecondary-process learning-and-memory based function invarious subcortical basal-ganglia to operate in ever moreprecise, life-supporting ways (Fig. 1). Those two evolutionaryfoundations may permit tertiary (neocortical) levels of brain-mind organization that indeed allows, recursively, for thekind of “reflective/recursive cognitive consciousness” thatenables and allows for higher-order mentation that Rollsskillfully focuses upon.

1. Evolutionary levels of control in brain andmind versus behavioristic biases

My bottom-up view contrasts starkly with Rolls’s top-downvision of consciousness, where the experience of qualia re-quires Higher Order Syntactic Thoughts (HOSTS). The two canbe synthesized, but for now, Rolls seems exceedingly defini-tive about some indefinite issues, by asserting that organismscannot have raw conscious experiences (qualia) withoutsyntactic functions comparable to human language [e.g., pp.492 where Rolls indicates that his “account implies that .

animals with a higher-order belief and thought system andwith linguistic (i.e., syntactic, not necessarily verbal) symbolmanipulation that have qualia.”]. I think that view, importantas it is for higher order thoughts (HOTs) ignores abundantevidence to the contrary, especially with respect to the primalaffective (raw-feeling) aspects of emotional, homeostatic andsensory arousals.

Although language is surely critical for high-level human-types propositional-cognitive consciousness, there is no hardevidence that raw affective experiencesdfrom physical andemotional “pains” to sensory and emotional delightsdrequirehigh-level syntactic (cognitive recursive-reflective) functionsin order to be experienced. Abundant data, largely from DeepBrain Stimulation (DBS) studies, in both humans and ‘lower’animals, evaluating the rewarding and punishing propertiesof such state shifts (perhaps instantiated in broad scale neu-rodynamics, perhaps attractor landscapes),which do not rely onneocortical processing (e.g., Huston & Borbely, 1973, 1974;Valenstein, 1966), are consistent with the conclusion thataffective-valuative coding is a rather primordial function ofmammalian brains (evolutionarily bright but cognitivelydull)dan experiential aspect of subcortical networks thatprovide distinct survival advantages, which may help explainwhy taste value is coded lower in the brains of rats than pri-mates, as Rolls’s research has demonstrated. Again,emotional feelings in animals are inferred from such DBSstudies by evaluating the rewarding and punishing propertiesof such artificially induced shifts in animals’ internal states (acritical issue not emphasized by Rolls), along with affectiveshifts that routinely transpire in human provoked by similartypes of DBS (Panksepp, 1985). I expect Rolls might suggestsuch rewarding and punishing brain effects are implicit andunconscious, based onmany humanmasking studies he cites,but hardly any of those studies evaluated, with the mostsensitive tools, changes in experienced affect, as we did inhumans, and surprisingly, observed reliable affective shiftsprovoked by cognitive information (emotional words) pre-sented tachistoscopically at 1 msec, substantially belowhuman visual absolute detection limits (Shevrin, Panksepp,Brakel, & Snodgrass, 2012).

As a result of our different perspectives on evolutionary“layering” of consciousnessdwith affective consciousnesspreceding cognitive variants (from raw feelings, to the brain’scapacity for HOTs to HOSTs, so to speak), we apparentlydisagree profoundly on the evolved reality of affective qualia,especially emotional feelings in other animals. For Rolls“reinforcement” seems to be a strictly unconscious process,whereas for me reinforcement is guided by brain valuative

Fig. 1 e Conceptual depiction of affective processing in thebrain: nested hierarchies of psychobehavioral control thehierarchical bottom-up and top-down (circular) causationthat is proposed to operate in every primal affective systemof the brain (with the SEEKING “enthusiasm” system beingdepicted, with different nomenclatures at three respectivelevels. It is proposed that in order for higher MindBrainfunctions to mature and function (via bottom-up control),they have to be integrated with the lower BrainMindfunctions, with primary affective processes being depictedas squares (SEEKING level), secondary-process learning ascircles (“wanting” level of analysis), and tertiary processes(“reward prediction level of analysis”) by rectangles. Thisaims to convey the manner in which bottom-up evolutionof nested hierarchies can integrate lower brain functionswith higher brain functions (in ways consistent withRolls’s HOST hypothesis) to eventually exert top-downregulatory control. Bottom-up affective controls, withcharacteristic emotive action patterns, prevail in early-infancy and early-childhood development. Top-downcontrol is optimized in late adolescence and adulthood,leading developmentally to diverse negatively andpositively valenced ruminations and daydreams (wishfulmind-wandering), respectively.

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circuits that elaborate raw affective qualiadthe various un-conditionally rewarding and punishing (i.e., valenced)subcortical network activities of the brain. It is noteworthythat DBS of the neocortex (which is essential for HOSTs) hasno such rewarding or punishing properties. I envision large-scale subcortical neurodynamic affective shifts in animalbrains to be a critical dimension of “reinforcement”, eventhough such “affective consciousness” (constituted of rawvaluative qualia) is not yet self-reflective, in ways Rollssuperbly envisions human decision-making to be. Rolls (inline with behaviorist traditions) sees no logical nor empiricalreasons to entertain the likelihood that unconditionalemotional arousals are experienced by other animals [e.g., thefreezing and flight evoked by foot shock, is not experienced byrats, despite all outward appearances and the fact that DBS ofsuch FEAR systems is punishing (Panksepp, et al., 1991)].

While I understand that radical behaviorism dealt veryeffectively with the outputs of the secondary learning andmemory processes of the brain (see Fig. 1), with “fear condi-tioning” being the greatest success story of behavioralneuroscience (LeDoux, 2012)dwork conducted by in-vestigators who choose to ignore the potential affectivequalities of primary-process emotional, homeostatic andsensory arousals because those issues seemed empiricallyimpenetrable in the pre-neuroscience era. Behavioristic“never-mind” skepticisms were epistemologically under-standable before the middle of the 20th century, since theascription of experiential states to non-speaking animalswould have been a gratuitous gesture until neuroscience dis-coveries indicating the rewarding and punishing properties ofdiencephalic DBS in animals (effects that survive radicalneodecortication of rats). As soon aswe could directlymonitorthe rewarding and punishing properties of specific brain net-works, a rich discussion about the psychological meaning ofsuch effects should have ensued, but the conversation hasbeen constrained if not outright discouraged/prohibited tillthe present day. I suspect a great opportunity to enrich ourunderstanding of subjectively experienced intrinsic brainfunctions was missed.

In this context, it should be recalled that behaviorism did aremarkably poor job of characterizing the sources andefficacy-attributes of the unconditioned stimuli (UCSs) andunconditioned responses (UCRs)dthe primary processes thatare critically important for learning to occur (for that wouldhave required neuroscience). It was a reasonable gambit toassume such brain processes are deeply unconscious beforeneuroscience. However, by the time neuroscience matured(the first meeting of the Society for Neuroscience was in 1972),the prohibition against going to the subjective level in otheranimals was sustained consensually within the newlymintedbehavioral neuroscience community, so no rich conversationabout the nature of subjective experiences ensued, evenwhenit became clear that wherever in the brain one could evokecoherent emotional patterns with DBS, those states wereconsistently rewarding or punishing. This is about as close toa lawful functional relationship as one has in brain research,and on the surface, it strongly suggests certain affects werebuilt into the brain by evolution. Simply having the word“reinforcement” is not a negation of that idea. It would nothave been far-fetched to posit that reinforcement was, in fact,

constituted from neural circuits controlling shifts in affectivelyexperienced states. Perhaps a form of “affective conscious-ness” preceded the evolution of a cognitive consciousness thateventually allowed us to have HOTs.

2. The subcortical sources of affective qualia

The implications of an “affective consciousness” existing inother animals, relying heavily on valenced subcortical net-works, might have been enormous for the development of ascientific psychology, but there was no institutionalconsensus that this was a reasonable path to take. Rolls barelymentioned this alternative perspective on affective qualia.Still, it remains possible, I believe likely, that an understand-ing of such intrinsic, low-level (primary-process) valuativebrain processes is critical for fathoming all other forms ofconsciousness permitted by higher brain-mind evolution, thatRolls discusses with grace and clarity.

In any event, a compelling empirical case can be made forthe provisional conclusion that raw emotional feelings arisefrom instinctual emotional-action neurodynamics, yieldingraw psychological states (affects) that predict survival tra-jectoriesdfor instance, unconditional appetitive (SEEKING)approach (for “enthusiastic” pursuit of resources), FEARfulwithdrawal (for feelings of “anxiety”), and explosiveenRAGEment agonistic engagements (for understanding raw“anger”). I use capitalizations routinely to distinguish suchprimary processes, to avoid mereological fallacies. These rawstates of mind are surely elaborated at higher lev-elsdengendering affect laden HOSTs, commonly calledruminationsdin recursive ways proposed by Rolls, but atpresent, there are no coherent lines of evidence that uncon-ditioned affective-emotional valuations emerge from higherneocortical processes, in any species (although Rolls hasshown that ancient orbitofrontal areas do participate in tasterelated hedonic shifts). In contrast, abundant lines of evidenceindicate that unconditional emotional rewarding and pun-ishing feelings are closely linked to brain networks thatmediate emotional action tendencies arising from very lowand ancient neural networks of mammalian brains, poten-tially constituting the first rudiments of “mind” that existedon the face of the earth (Panksepp, 1982, 1985, 1998; Panksepp& Biven, 2011). The view I have proposed has not beennegated.

Thus, I would urge Rolls to reconsider the likelihood thatprimary-process neural circuits for certain unconditionedsubcortical emotional, homeostatic and sensory responses,are critical in constituting raw affective qualia. Hunger andanger may be experienced by rats within subcortical brainregions. Neglect of such processes may delay medicinal de-velopments (Panksepp, 2010; Panksepp, Wright, Dobrossy,Schlaepfer, & Coenen, 2014). Why might such unreflectivestates of mind exist? A reasonable evolutionary answer is thatthey were incredibly useful for survival and reproduction: Inother words, the function of raw affective feelings is tointrinsically anticipate survival issues, and thereby to promotelearning and memory, to further maximize opportunities forsurvival with increasingly sophisticated behavioral andcognitive strategies. In this view, primal sensations of “pain”

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are useful since they intrinsically predict survival trajectories,etc. etc. This is not to deny the important role of affectiveHOSTs, sowell developed by Rolls, in highermental processes.

To reiterate, robust evidence for the existence of variousaffective states in other animal is simple: For over 60 years wehave known fromDBS studies that direct arousal of emotionalcircuits can reinforce behavioral changesdespecially varioussimple approach and avoidance behaviors. There is abundantevidence that artificial arousals of such rewarding and pun-ishing subcortical emotional circuits feel like something .

most explicitly in humans (Panksepp, 1985). For instancewhen the dorsal Periaqueductal Gray (PAG) trajectory of theFEAR system was first stimulated, a patient suddenlyexclaimed being “scared to death”! (Nashold, Wilson, &Slaughter, 1969). Many such affective responses seem toofast for HOSTs to be recruiteddjust think of the suddenness ofsharp pain. There are emerging clinical examples, with themost recent spectacular finding being based on DBS of the so-called “brain reward system”dthe massive Medial ForebrainBundle based SEEKING network (Panksepp, 1982, 1998)dwhich can produce remarkable anti-depressant effects inhuman (Schlaepfer, Bewernick, Kayser, Madler, & Coenen,2013). And this is not because the system engenders feelingsof sensory hedonics, as many behavioral investigators origi-nally entertained (e.g., Berridge & Valenstein, 1991; Wise,1982), but states of positive enthusiasm, that normallyaccompany the appetitive-foraging phase of behavior in ourspecies, as we have long predicted (from Panksepp, 1981, 1982to Coenen, Schlaepfer, Maedler, & Panksepp, 2011, 2012). Whythis systemwas simply labeled “the brain reward system” andlargely left at that, reflects how rigidly radical behaviorismhasguided the conversation till now, with abundantconsensually-sustained constraints to never stray into theexperiential realm in other animals.

It is certainly not an illogical step to surmise that compa-rable feelings arise in non-speaking animals. There is abun-dant reason to believe that feelings associated with such rawemotional arousals are not simply higher brain “read-outs” oflower brain processes (Damasio, 2010; Panksepp, 1981, 1998,2005) but actually constituted the first glimmers of con-sciousness (qualia) that existed on the face of the earth. If so,we would be wise to prioritize the role of neural circuits thatmediate raw affects in the constitution of learning and higherlevels of consciousness (an idea barely discussed by Rolls, aswell as the consciousness literature at large), rather than tostill believe in some kind of amorphous non-sentient conceptsof ‘rewards’ and ‘punishments’ passed down to us from a pre-neuroscientific behaviorist era that had no access to norapparent interest in abundant neuroscientific evidence foraffective states in animals.

Parenthetically, it is worth noting that even meta-analysesof available brain Functional Magnetic Resonance Imaging(fMRI)-imaging studies suggest humans also have comparableemotional feelings (Lench, et al., 2011, 2013; Vytal & Hamann,2010, albeit vigorously opposed by Lindquist, Wager, Kober,Bliss-Moreau, & Barrett, 2012, 2013), with deep subcorticalloci of control, as measured by the most well-conceived brainPositron Emission Tomography (PET)-imaging studies ofhuman emotional feelings (e.g., Damasio, et al., 2000, and forcomparable homeostatic work see Denton, 2006). There is

inadequate space here to cover such human brain imagingresearch, but I simply note that fMRI studies probably are notwell suited for imaging affective processes, since we cannotcontrol timing-analytical issues properly, while PET studiesare muchmore suited for that task. Indeed, the best studies inthat genera highlight deep subcortical arousals as being crit-ical for basic human emotional experiences, namely fear,anger, sadness and joy (Damasio, et al., 2000), and with fMRI,affective changes are positively correlated to subcorticalarousal rather than cortical ones with fMRI (Northoff, et al.,2009), but only as long as one uses methods that avoid con-founds from cognitive-emotional reciprocal interactions.Such effects are opposite to those predicted by Rolls’s syn-tactic HOST theory, where emotional feelings should bepositively correlated to higher brain arousals.

3. Thus, where in the brain are rewardsrewarding and punishers punishing?

I would judiciously ask Rolls to clarify the logical problems hefinds with an affective neuroscience approach which relies onaffect being empirically indexed by the rewarding and pun-ishing properties of direct electrical stimulation of brainemotional circuits (Panksepp, 1982, 1998). Of course one couldpostulate that such rewarding and punishing states are notexperienced in animals or we have no way of knowing, butwhen appropriate scientific skepticism is countered by “theweight of evidence”dwith animal research well predictinghuman affective responses (Panksepp, 1985)dwemay be wiseto deal with the converging evidence for the most parsimo-nious interpretation, namely that primal rewards and pun-ishments in animals are affectively experienced withinsubcortical networks. For that not to be the case, someone(Rolls?) needs to explain, with clear predictions, how “re-wards” and “punishments” could operate totally uncon-sciously, with no experiences transpiring in animal brains,while evolutionarily similar (homologous?) states of mind areexperienced in human brains. HOST theory may be able to dothat, but perhaps currently only by stipulation rather thandifferential scientific predictions. Parenthetically, Rolls and Iagree that cortical-readout theories, such as the famousJames-Lange conjecture, remain largely without neuroscien-tific substance as major sources of primal affective feelings.Still, considering that syntactic brain functions probablyrequire cortex, Rolls should predict that most affective shiftsemerge from cortex, and to my knowledge there is no evi-dence for that conjecture. If not, we scientists must abide bythe existing weight of evidence.

Overall, there is nothing in Rolls’s data or theorizing toconvince me that distinct ‘qualia-tative’ forms of “phenom-enal affective feelings” emerged in brain evolution only withthe emergence of cortically-mediated syntactic thought pro-cesses. I think that provided the capacity for explicit cognitivetestimony of our experiences. Despite a wonderfully rich bookon the brain, Rolls still has the job of empirically demonstratingthat (and then how) neocortexdwhich clearly is not needed fordramatic emotional displays, such as decorticate rage nor forbrain “reward” or “punishment” ddeveloped any intrinsiccapacity to mediate positive and negative affective feelings.

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How did neocortex grant us the opportunity to experienceintense affective states? Everyone would surely agree thatneocortex allows us to conceptualize ourselves as feelingcreatures, and also provides essential neural substrates formaturation of complex cognitive lives. But where is the evi-dence that our expansive neocortex allows us to feel thepleasures and pains of existence, especially when neonatallyneodecorticated animals and humans exhibit diverse basicemotions andmotivations (Solms & Panksepp, 2012), even thejoy of social play (Panksepp, Normansell, Cox, & Siviy, 1994)?Understandably many behavioral scientists have been skep-tical of such psychological discoveries in animals (for instancejust a few years ago LeDoux (2012, p. 666) asserted “We willnever know what an animal feels.” Of course, our cardinalvalue of infinite skepticism always needs to bow to the“weight of evidence”. Thus, in order for science to flourish, ourskepticism also needs to be tempered by our appreciation ofthe remaining cardinal value of scientific thinking, which isthat in science there is no absolute knowledge. That shoulddiscourage us from making premature “never” statements.

I would submit that at the present moment the “weight ofevidence” strongly supports the conclusion that all mammalsexperience their emotional arousals andhomeostatic ones also(e.g., hunger and thirst), and that the critical neural mecha-nisms are subcortically concentrated, with abundant (HOSTsof) effects on neocortical cognitive functions. Such primal af-fects are beginning to be studied neuroscientifically in animalmodels, which I expect will promote effective medicinaldevelopment for many psychiatric disorders and other affec-tive problems. Whether animals cognitively reflect upon theiraffectively experiences remains scientifically unpenetrated,and so far Rolls’s beautifully crafted HOST theory, which Iadmired greatly, applies well mostly to human decision-making processes. Although Rolls has collected some wonder-fully suggestive correlative data for animal “cognitions” (e.g., seepp. 406e409 of his book), I feel confident that Rolls would agreethat the constitution of such higher mental processes remainremarkably resistant to neuroscientific analysis, especially inotheranimals. Incontrast, I suspectaffectiveneuroscience typeanimal-modeling of human emotional feeling will give us solidneuroscientific insights into human psychiatric problems (e.g.,Panksepp, 2005, 2006; Panksepp et al., 2014).

4. Implications of affective processes inanimal for human therapeutics

A scientific analysis of the basic affective processes ofmammalianbrainsmayalsohaveprofound implication for theway we eventually understand and treat other animals, andperhaps each other. In my estimation, Rolls’s fascinatingHOSTs theory for higher mental processes, by suggesting itmay also explain affective qualia (e.g., pp. 491e493), comesrather lose to suggesting thatmanyof the othermammals, likethose ubiquitous laboratory rats andmicewe effectively use tostudy behavior, have no feelings (minds?) at alldthat they arecloser tozombies thansentient-feelingcreatures. I think that ismost unlikely, based on many DBS (causal) studies of howemotional feelings emerge from subcortical circuits. In thiscontext, I would note that our discovery of “rat laughter” and

the fact that those tickle-induced sounds, arise from braincircuits that rats find remarkably rewarding (Burgdorf, Wood,Kroes, Moskal, & Panksepp, 2007), in an “enthusiasm” pro-moting “brain reward-SEEKING system” way, has opened upthe development of several new anti-depressant concepts(Burgdorf, Panksepp, & Moskal, 2011; Panksepp, et al., 2014)that are currently in human testing, with positive results.

In sum, I suspect we will never understand how primalaffective statesdthat surely help constitute some of ourfundamental psychological valuesdarise from neural activ-ities without incisive animal models. It would seem thatRolls’s theory, that affective qualia in humans arise fromHOSTs, would make that a fool’s errand. So far our novel,cross-species affective neuroscience-guided, discoveries inanti-depressant development are inconsistent with that like-lihood (Burgdorf, et al., 2011; Panksepp, et al., 2014). PerhapsRolls’s remarkably sophisticated analysis of decision-makingwould benefit from more explicit bottom-up conceptualiza-tions of the evolution of the affective mind, with uncondi-tional brain primary affective process circuitsdemotional,homeostatic and sensorydcontrolling secondary-processes oflearning and memory (the brain mechanisms of associativelearning probably being deeply unconscious)dwhich provideraw psychological materials that require sophisticated deci-sion making to further facilitate thriving (Narvaez, Panksepp,Schore, & Gleason, 2013).

5. Concluding reflections

This said, Rolls’s Emotion and Decision Making book is a treasuretrove of important information that has to be integrated into acoherent whole if we are to understand the kinds of creaturesthat we really are. His recent book is a must-read for anyonewho has wondered about the brain-processes that make uspsychologically alive. Whether neuroscientific understandingand mathematical modeling of our decision-making abilitieswill get us closer to understanding various affective dynamicsthat routinely control our thinking processes remains debat-able. Still, a true neural understanding of affective states willbe of momentous importance for understanding humanconsciousness and promoting a scientific psychiatry. I thinkwe have learned a great deal about human affects by studyinghomologous emotional processes in other animals (Panksepp& Biven, 2011; Panksepp et al., 2014). I look forward to the daywhen HOST-guided research into human emotions furtherfacilitates our understanding of the neural channeling of bothproductive thoughts as well as disturbing ruminations. So farwe can be confident that such higher mental processes will,forever, remain much, MUCH easier to study in humans thanin other animals. On this, I expect Edmund and I willcompletely agree.

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Received 23 April 2014Accepted 23 April 2014

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Please cite this article in press as: Panksepp, J., Integrating bottom-up internalist views of emotional feelings with top-downexternalist views: Might brain affective changes constitute reward and punishment effects within animal brains?, Cortex(2014), http://dx.doi.org/10.1016/j.cortex.2014.04.015